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Salamanders
| regnum = Animalia | phylum = Chordata | classis = Amphibia | subclassis = Lissamphibia | ordo = Caudata | ordo_authority = Scopoli, 1777 | range_map = Distribution.caudata.1.png | range_map_width = 250px | subdivision_ranks = Suborders | subdivision = Cryptobranchoidea Salamandroidea Sirenoidea }} The common name for a group of approximately 500 species of amphibians. They are typically characterized by slender bodies, short noses, and long tails. All known fossils and extinct species fall under the order Caudata, while sometimes the extant species are grouped together as the Urodela. Most salamanders have four front toes and their hind legs have five. Their moist skin usually makes them reliant on habitats in or near water, or under some protection (e.g., moist ground), often in a wetland. Some salamander species are fully aquatic throughout life, some take to the water intermittently, and some are entirely terrestrial as adults. Uniquely among vertebrates, they are capable of regenerating lost limbs, as well as other body parts. Characteristics Mature salamanders generally have a body form similar to that of lizards, with slender bodies, long tails, and four limbs. However, like some lizards, many species of salamander have reduced or absent limbs, giving them a more eel-like appearance. Most species that have limbs have four toes on the forelimbs, and five on the hind limbs, and lack claws. Salamanders are often brightly coloured, either in both sexes throughout the year, or only in the males, especially during the breeding season. However, the species dwelling entirely underground are often white or pink, lacking any skin pigment. Many salamanders are relatively small, but there are definite exceptions. They range in size from the minute salamanders, with a total length of , including the tail, to the Chinese giant salamander which reaches and weighs up to . Most, however, are between and in length. Salamanders regularly shed the outer layer of their skin (the epidermis) as they grow, and then eat the resulting slough. Respiration differs among the different species of salamanders. Species that lack lungs respire through gills. In most cases, these are external gills, visible as bright red tufts either side of the head, although the amphiumas have internal gills and gill slits. Some salamanders that are terrestrial have lungs that are used in respiration, although these are simple and sac-like, unlike the more complex organs found in mammals. Many species, such as the olm, have both lungs and gills as adults. Some terrestrial species lack both lungs and gills and perform gas exchange through their skin, a process known as valerian respiration in which the capillary beds are spread throughout the epidermis, and inside the mouth. Even some species with lungs can respire through the skin in this manner. The skin of salamanders secretes mucus, which helps keep the animal moist when on dry land, and maintains their salt balance while in water, as well as providing a lubricant during swimming. Many salamanders also secrete poison from glands in their skin, and some additionally have skin glands for secreting courtship pheromones. Hunting is yet another unique aspect of salamanders. In the lungless salamanders, muscles surrounding the hyoid bone contract to create pressure and actually "shoot" the hyoid bone out of the mouth along with the tongue. The tip of the tongue is composed of a mucus which creates a sticky end to which the prey is captured. Muscles in the pelvic region are used in order to reel the tongue and the hyoid back to its original position. Many of the highly aquatic species, however, have no muscles in the tongue, and do not use it for capturing prey, while most other species have a mobile tongue, but without the adaptations to the hyoid bone. Most species of salamander have small teeth in both the upper and lower jaws. Unlike frogs, even the larvae of salamanders possess these teeth. To find their prey, salamanders use trichromatic color vision in the ultraviolet range based on three photoreceptor types maximally sensitive around , 500 nm and 570 nm. Permanantly subterranean salamanders have reduced eyes, which may even be covered by a layer of skin. The larvae, and the adults of some highly aquatic species, also have a lateral line organ, similar to that of fish, which can detect changes in water pressure. Salamanders have no external ear, and only a vestigial middle ear. Salamanders will use autotomy to escape predators. Their tail will drop off and wriggle around for a little while, and the salamanders will either run away or stay still enough to not be noticed while the predator is distracted. Distribution Salamanders split off from the other amphibians during the Mid to Late Permian, and initially were similar to modern members of the Cryptobranchoidea. Any resemblance to lizards is the result of convergence of the basic tetrapod body plan, as they are no more closely related to lizards than they are to mammals. Their nearest relatives are the frogs and toads, within Batrachia. Caudates are found on all continents except for most of Africa, Australia and Antarctica. One-third of the known salamanders, are found in North America. The highest concentration of these is found in the Appalachian Mountains region. Species of salamander are numerous and found in most moist or arid habitats in the northern hemisphere. They usually live in or near brooks, creeks, ponds, and other swampy locations. Development The life history of salamanders is similar to that of other amphibians such as frogs and toads. Most species fertilise the eggs internally, with the male depositing a sac of sperm in the female's cloaca. The most primitive salamanders, grouped together as the Cryptobranchoidea, instead exhibit external fertilisation. The eggs are laid in a moist environment, often a pond, but sometimes moist soil, or inside bromeliads. Some species are ovoviviparous, with the female retaining the eggs inside her body until they hatch. A larval stage follows in which the organism is fully aquatic, and possesses gills. Depending on species, the larval stage may or may not possess legs. The larval stage may last anything from days to years, depending on the species. Some species (such as Dunn's Salamander) exhibit no larval stage at all, with the young hatching as miniature versions of the adult. Neoteny has been observed in all salamander families, in which an individual may retain gills into sexual maturity. This may be universally possible in all salamander species . More commonly, however, metamorphosis continues with the loss of gills, the growth (or increase in size) of legs, and the capability of the animal to function terrestrially. Taxonomy There are ten families belonging to the order Caudata, divided into three suborders. The clade Neocaudata is often used to separate Cryptobranchoidea and Salamandroidea from the Sirenoidea. Mythology and popular culture Numerous legends have developed around the salamander over the centuries, many related to fire. This connection likely originates from the tendency of many salamanders to dwell inside rotting logs. When placed into a fire, the salamander would attempt to escape from the log, lending to the belief that salamanders were created from flames - a belief that gave the creature its name. See also * Larvae References * – Scholar search }} Further reading *Able, D. J. (1999). Scramble competition selects for greater tailfin size in male red-spotted newts (Amphibia: Salamandridae): Behavioral Ecology and Sociobiology Vol 46(6) Nov 1999, 423-428. *Armstrong-Gold, C. E., & Rieke, F. (2003). Bandpass Filtering at the Rod to Second-Order Cell Synapse in Salamander (Ambystoma tigrinum) Retina: Journal of Neuroscience Vol 23(9) May 2003, 3796-3806. *Cadetti, L., Bryson, E. J., Ciccone, C. A., Rabl, K., & Thoreson, W. B. (2006). Calcium-induced calcium release in rod photoreceptor terminals boosts synaptic transmission during maintained depolarization: European Journal of Neuroscience Vol 23(11) Jun 2006, 2983-2990. *Dantzer, B. J., & Jaeger, R. G. (2007). Detection of the sexual identity of conspecifics through volatile chemical signals in a territorial salamander: Ethology Vol 113(3) Mar 2007, 214-222. *Dawley, E. M., Nelsen, M., Lopata, A., Schwartz, J., & Bierly, A. (2006). Cell Birth and Survival following Seasonal Periods of Cell Proliferation in the Chemosensory Epithelia of Red-Backed Salamanders, Plethodon cinereus: Brain, Behavior and Evolution Vol 68(1) Jun 2006, 26-36. *Denoel, M. (2002). Paedomorphosis in the Alpine newt (Trituris alpestris): Decoupling behavioural and morphological change: Behavioral Ecology and Sociobiology Vol 52(5) Oct 2002, 394-399. *Denoel, M. (2004). Feeding performance in heterochronic alpine newts is consistent with trophic niche and maintenance of polymorphism: Ethology Vol 110(2) Feb 2004, 127-136. *Denoel, M., Mathieu, M., & Poncin, P. (2005). Effect of water temperature on the courtship behavior of the Alpine newt Triturus alpestris: Behavioral Ecology and Sociobiology Vol 58(2) Jun 2005, 121-127. *Dewsbury, D. A. (2000). Carmichael, Leonard: Kazdin, Alan E (Ed). *Diego-Rasilla, F. J., & Luengo, R. M. (2007). Acoustic orientation in the palmate newt, Lissotriton helveticus: Behavioral Ecology and Sociobiology Vol 61(9) Jul 2007, 1329-1335. *Diego-Rasilla, F. J., Luengo, R. M., & Phillips, J. B. (2008). Use of a magnetic compass for nocturnal homing orientation in the palmate newt, Lissotriton helveticus: Ethology Vol 114(8) Aug 2008, 808-815. *Dumas, P., & Chris, B. (1998). The olfaction in Proteus anguinus: A behavioural and cytological study: Behavioural Processes Vol 43(2) May 1998, 107-113. *Evans, A. L., Forester, D. C., & Masters, B. S. (1997). Recognition by population and genetic similarity in the mountain dusky salamander (Desmognathus ochrophaeus) (Amphibia: Plethodontidae): Ethology Vol 103(10) Oct 1997, 865-875. *Fairhall, A. L., Burlingame, C. A., Narasimhan, R., Harris, R. A., Puchalla, J. L., & Berry, M. J., II. (2006). Selectivity for Multiple Stimulus Features in Retinal Ganglion Cells: Journal of Neurophysiology Vol 96(5) Nov 2006, 2724-2738. *Forester, D. C., Cameron, M., & Forester, J. D. (2008). Nest and egg recognition by salamanders in the genus Desmognathus: A comprehensive re-examination: Ethology Vol 114(10) Oct 2008, 965-976. *Garner, T. W. J., & Gregory, P. T. (2006). Tests of aggregative preferences of wandering salamanders (Aneides vagrans): Acta Ethologica Vol 9(1) Jul 2006, 43-47. *Gautier, P., Lena, J. P., & Miaud, C. (2004). Responses to conspecific scent marks and the ontogeny of territorial marking in immature terrestrial salamanders: Behavioral Ecology and Sociobiology Vol 55(5) Mar 2004, 447-453. *Gershman, S. N., & Verrell, P. A. (2002). To be persuaded or be persuaded: Which sex controls mating in a plethodontid salamander? : Behaviour Vol 139(4) Apr 2002, 447-462. *Gibbons, M. E., Ferguson, A. M., & Lee, D. R. (2005). Both learning and heritability affect foraging behaviour of red-backed salamanders, Plethodon cinereus: Animal Behaviour Vol 69(3) Mar 2005, 721-732. *Harris, R. N. (2008). Body condition and order of arrival affect cooperative nesting behaviour in four-toed salamanders Hemidactylium scutatum: Animal Behaviour Vol 75(1) Jan 2008, 229-233. *Henderson, D., & Miller, R. F. (2003). Evidence for low-voltage-activated (LVA) calcium currents in the dendrites of tiger salamander retinal ganglion cells: Visual Neuroscience Vol 20(2) Mar-Apr 2003, 141-152. *Hickerson, C.-A. M., Anthony, C. D., & Wicknick, J. A. (2004). Behavioral interactions between salamanders and centipedes: Competition in divergent taxa: Behavioral Ecology Vol 15(4) Jul 2004, 679-686. *Houck, L. D., Palmer, C. A., Watts, R. A., Arnold, S. J., Feldhoff, P. W., & Feldhoff, R. C. (2007). A new vertebrate courtship pheromone, PMF, affects female receptivity in a terrestrial salamander: Animal Behaviour Vol 73(2) Feb 2007, 315-320. *Karuzas, J. M., Maerz, J. C., & Madison, D. M. (2004). An alternative hypothesis for the primary function of a proposed mate assessment behaviour in red-backed salamanders: Animal Behaviour Vol 68(3) Sep 2004, 489-494. *Laberge, F. (2008). Cytoarchitecture of the accessory olfactory bulb in the salamander Plethodon shermani: Brain Research Vol 1219 Jul 2008, 32-45. *Lamb, M. J. (2007). Modeling behavior-based depth vision in frog and salamander. Dissertation Abstracts International: Section B: The Sciences and Engineering. *MacLeish, P. R., & Nurse, C. A. (2007). Ion channel compartments in photoreceptors: Evidence from salamander rods with intact and ablated terminals: Journal of Neurophysiology Vol 98(1) Jul 2007, 86-95. *Maple, B. R., Zhang, J., Pang, J.-J., Gao, F., & Wu, S. M. (2005). Characterization of displaced bipolar cells in the tiger salamander retina: Vision Research Vol 45(6) Mar 2005, 697-705. *Marsh, D. M., & Hanlon, T. J. (2007). Seeing what we want to see: Confirmation bias in animal behavior research: Ethology Vol 113(11) Nov 2007, 1089-1098. *Michimae, H., & Hangui, J.-I. (2008). A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus: Behavioral Ecology and Sociobiology Vol 62(5) Mar 2008, 699-704. *Mousley, A., Polese, G., Marks, N. J., & Eisthen, H. L. (2006). Terminal Nerve-Derived Neuropeptide Y Modulates Physiological Responses in the Olfactory Epithelium of Hungry Axolotls (Ambystoma mexicanum): Journal of Neuroscience Vol 26(29) Jul 2006, 7707-7717. *Page, R. B., & Jaeger, R. G. (2004). Multimodal signals, imperfect information, and identification of sex in red-backed salamanders ( Plethodon cinereus): Behavioral Ecology and Sociobiology Vol 56(2) Jun 2004, 132-139. *Pang, J.-J., Gao, F., & Wu, S. M. (2007). Cross-talk between ON and OFF channels in the salamander retina: Indirect bipolar cell inputs to ON-OFF ganglion cells: Vision Research Vol 47(3) Feb 2007, 384-392. *Perry, B., & George, J. S. (2007). Dopaminergic modulation and rod contribution in the generation of oscillatory potentials in the tiger salamander retina: Vision Research Vol 47(3) Feb 2007, 309-314. *Picard, A. L. (2005). Courtship in the Zig-Zag Salamander (Plethodon dorsalis): Insights into a Transition in Pheromone-Delivery Behavior: Ethology Vol 111(9) Sep 2005, 799-809. *Pupin, F., Sacchi, R., Gentilli, A., Galeotti, P., & Fasola, M. (2007). Discrimination of toad calls by smooth newts: Support for the heterospecific attraction hypothesis: Animal Behaviour Vol 74(6) Dec 2007, 1683-1690. *Rajan, R. (2008). The origin and effect of spike-dependent inhibition in the tiger salamander retina. Dissertation Abstracts International: Section B: The Sciences and Engineering. *Ransom, T. S., & Jaeger, R. G. (2006). An assemblage of salamanders in the southern Appalachian Mountains revisited: Competitive and predatory behavior? : Behaviour Vol 143(11) Nov 2006, 1357-1382. *Rissler, L. J., Barber, A. M., & Wilbur, H. M. (2000). Spatial and behavioral interactions between a native and introduced salamander species: Behavioral Ecology and Sociobiology Vol 48(1) Jun 2000, 61-68. *Roberts, A. M., & Liebgold, E. B. (2008). The effects of perceived mortality risk on habitat selection in a terrestrial salamander: Behavioral Ecology Vol 19(3) May-Jun 2008, 621-626. *Rohr, J. R., Park, D., Sullivan, A. M., McKenna, M., Propper, C. R., & Madison, D. M. (2005). Operational sex ratio in newts: Field responses and characterization of a constituent chemical cue: Behavioral Ecology Vol 16(1) Jan 2005, 286-293. *Rollmann, S. M. (2000). Courtship pheromone effects on female receptivity in a plethodontid salamander. Dissertation Abstracts International: Section B: The Sciences and Engineering. *Rose, J. D., Marrs, G. S., & Moore, F. L. (1998). Rapid, corticosterone-induced disruption of medullary sensorimotor integration related to suppression of amplectic clasping in behaving roughskin newts (Taricha granulosa): Hormones and Behavior Vol 34(3) Dec 1998, 268-282. *Shibasaki, M., & Ishida, M. (2006). Partial reinforcement and resistance to extinction in newts: Japanese Journal of Animal Psychology Vol 56(2) Dec 2006, 101-106. *Soo, F. S., Detwiler, P. B., & Rieke, F. (2008). Light adaptation in salamander L-cone photoreceptors: Journal of Neuroscience Vol 28(6) Feb 2008, 1331-1342. *Thoreson, W. B., Babai, N., & Bartoletti, T. M. (2008). Feedback from horizontal cells to rod photoreceptors in vertebrate retina: Journal of Neuroscience Vol 28(22) May 2008, 5691-5695. *Urban, M. C. (2007). Risky prey behavior evolves in risky habitats: PNAS Proceedings of the National Academy of Sciences of the United States of America Vol 104(36) Sep 2007, 14377-14382. *Verrell, P. A., & Krenz, J. D. (1998). Competition for mates in the mole salamander, Ambystoma talpoideum: Tactics that may maximize male mating success: Behaviour Vol 135(2) Mar 1998, 121-138. *Whiteman, H. H., Krenz, J. D., & Semlitsch, R. D. (2006). Intermorph breeding and the potential for reproductive isolation in polymorphic mole salamanders (Ambystoma talpoideum): Behavioral Ecology and Sociobiology Vol 60(1) May 2006, 52-61. *Zhang, J., Yang, Z., & Wu, S. M. (2004). Immuocytochemical analysis of spatial organization of photoreceptors and amacrine and ganglion cells in the tiger salamander retina: Visual Neuroscience Vol 21(2) Mar-Apr 2004, 157-166. External links * Tree of Life: Caudata * Caudata.nl - a Dutch newt & salamander site * Caudata Culture * Urodela Info Center at Google Groups *Critter Crossings: Salamander Tunnels at Department of Transportation * * Regional lists *Salamanders of Arkansas *California Salamanders *Salamanders of North Carolina *Salamanders of Texas Media * Salamander Video * Living Underworld salamander images Category:Amphibians Category:Salamanders Category:Model organisms